Australian Journal of Botany Australian Journal of Botany Society
Southern hemisphere botanical ecosystems

A classification and census of regenerative strategies in the eucalypts (Angophora, Corymbia and Eucalyptus—Myrtaceae), with special reference to the obligate seeders

Dean Nicolle
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Currency Creek Arboretum Eucalypt Research, 15 Rousillion Promenade, Old Reynella, SA 5161, Australia. Email:

Australian Journal of Botany 54(4) 391-407
Submitted: 31 March 2005  Accepted: 31 October 2005   Published: 22 June 2006


A survey of regenerative strategies in the eucalypts, including lignotuber development, was undertaken by extensive field observations, seedling trials and trials of cultivated individuals over a 12-year period. Four broad regenerative strategies were identified, viz. obligate seeders, lignotuber sprouters, stem sprouters and combination sprouters. These four regenerative strategies are based on the ability to develop a lignotuber and the regeneration strategy after whole-crown destruction. These regenerative strategies do not wholly correspond to the tree, mallee, mallet, marlock and shrub habit categories commonly applied to eucalypts. The obligate seeders include many more terminal taxa than have been previously documented as mallet taxa, with 78 western obligate seeders (the ‘true’ mallets) and nine eastern obligate seeders listed herein. Obligate seeders do not possess a lignotuber and are killed by crown-destructive events, and as such are relatively short-lived in most natural environments. A further 16 taxa are also known to be non-lignotuberous, but these are capable of producing epicormic regrowth from the trunk following crown destruction and are defined as stem sprouters. The remaining two regenerative strategies include taxa that are both lignotuberous and able to regenerate vegetatively following disturbance events. The persistent and conjecturous mallet–marlock–moort distinction is rejected, this study showing it to be dependent on stand density. Data presented indicate no significant difference in germination time or maturation time between western obligate-seeder taxa and closely related sprouter taxa. The conservation status of obligate-seeder taxa is discussed. Nomenclatural issues regarding the taxonomic distinction between obligate-seeder and resprouter sister taxa are discussed. A census of regenerative strategies for all recognised eucalypt taxa is included as an accessory publication on the web.


This paper would not have been possible without the early encouragement of John Connors and encouragement and assistance of Malcolm Gill. I thank the many people who assisted in reading and commenting on earlier manuscripts of the paper, in particular Malcolm Gill, Malcolm French, Ian Brooker, Peter White, Stephen Hopper and Colin Yates.


Ashton DH (1981) Fire in tall open-forests (wet sclerophyll forests). In ‘Fire and the Australian biota’. (Eds AM Gill, RH Groves, IR Noble) pp. 243–272. (Australian Academy of Sciences: Canberra)

Atkins K (1998) Threatened flora legislation and listings. In ‘Western Australia’s threatened flora’. (Eds A Brown, C Thomson-Dans, N Marchant) pp. 21–23. (Department of Conservation and Land Management: Perth)

Barker WR , Barker RM , Haegi L (1999) Introduction to Hakea. In ‘Flora of Australia. Vol. 17B. Proteaceae. 3. Hakea to Dryandra’. (Ed. A. Wilson) pp. 1–30. (ABRS: Canberra/CSIRO: Melbourne)

Beard JS (1975) ‘Vegetation survey of Western Australia—Nullarbor.’ (University of Western Australia Press: Perth)

Beard JS (1990) The mallee lands of Western Australia. In ‘The mallee lands: a conservation perspective’. (Eds JC Noble, PJ Joss, GK Jones) pp. 29–33. (CSIRO: Melbourne)

Bennett EM (1995) Hybrid between Eucalyptus tetraptera and Eucalyptus stoatei from Jerdacuttup, Western Australia. Nuytsia 10(1), 1–5.

Blake TJ (1972) Studies on the lignotubers of Eucalyptus obliqua L’Herit. New Phytologist 71, 327–334.
CrossRef |

Boland DJ , Brooker MIH , Turnbull JW , Kleinig DA (1980) ‘Eucalyptus seed.’ (CSIRO: Melbourne)

Boland DJ , Brooker MIH , Chippendale GM , Hall N , Hyland BPM , Johnston RD , Kleinig DA , Turner JD (1984) ‘Forest trees of Australia.’ 4th edn. (CSIRO: Melbourne)

Briggs JD , Leigh JH (1996) ‘Rare or threatened Australian plants.’ (CSIRO Publishing: Melbourne)

Brooker MIH (2000) A new classification of the genus Eucalyptus L’Her. (Myrtaceae). Australian Systematic Botany 13, 79–148.
CrossRef |

Brooker MIH, Hopper SD (1991) A taxonomic revision of Eucalyptus wandoo, E. redunca and allied species (Eucalyptus series Levispermae Maiden—Myrtaceae) in Western Australia. Nuytsia 8(1), 1–189.

Brooker MIH, Hopper SD (1993) New series, subseries, species and subspecies of Eucalyptus (Myrtaceae) from Western Australia and from South Australia. Nuytsia 9(1), 1–68.

Brooker MIH, Hopper SD (2002) Taxonomy of species deriving from the publication of Eucalyptus subseries Cornutae (Myrtaceae). Nuytsia 14(3), 325–360.

Brooker MIH , Kleinig DA (1990) ‘Field guide to eucalypts. Vol. 2. South-west and southern Australia.’ (Inkata Press: Melbourne)

Burgess IP, Bell JC (1983) Comparative morphology and allozyme frequencies of Eucalyptus grandis Hill ex Maiden and E. saligna Sm. Australian Forest Research 13, 133–149.

Carnahan JA (1976) Natural vegetation. In ‘Atlas of Australian resources’. (Commonwealth of Australia, Department of Natural Resources: Canberra)

Carr DJ, Carr SGM (1980) The Lehmannianae: a natural group of Western Australian eucalypts. Australian Journal of Botany 28, 525–550.

Carr DJ, Carr SGM, Jahnke R (1982) The eucalypt lignotuber: a position-dependant organ. Annals of Botany 50, 481–489.

Carr DJ, Jahnke R, Carr SGM (1983) Development of the lignotuber and plant form in Lehmannianae.  Australian Journal of Botany 31, 629–643.
CrossRef |

Chippendale GM (1988) Eucalyptus, Angophora. In ‘Flora of Australia. Vol. 19. Myrtaceae—Eucalyptus, Angophora’. (Ed. AS George) pp. 1–456. (Australian Government Publishing Service: Canberra)

Cremer KW , Cromer RN , Florence RG (1984) Stand establishment. In ‘Eucalypts for wood production’. (Eds WE Hillis, AG Brown) pp. 81–135. (CSIRO and Academic Press: Sydney)

Eldridge KG (1963) Eucalyptus sieberiana × regnans—An artificial hybrid. Australian Forestry 27(2), 128–129.

Gardner CA (1957) The fire factor in relation to the vegetation of Western Australia. Western Australian Naturalist 5, 166–173.

George AS (1981) The genus Banksia L.f. (Proteaceae). Nuytsia 3(3), 239–473.

Gilbert JM (1959) Forest succession in the Florentine Valley, Tasmania. Proceedings of the Royal Society of Tasmania 93, 129–151.

Gill AM (1981) Adaptive responses of Australian vascular plant species to fires. In ‘Fire and the Australian biota’. (Eds AM Gill, RH Groves, IR Noble) pp. 243–272. (Australian Academy of Science: Canberra)

Gill AM (1997) Eucalypts and fires: interdependent or independent? In ‘Eucalypt ecology: individuals to ecosystems’. (Eds JE Williams, JCZ Woinarski) pp. 151–167. (Cambridge University Press: Cambridge, UK)

Gill AM, Ashton DH (1968) The role of bark type in relative tolerance to fire of three central Victorian eucalypts. Australian Journal of Botany 16, 491–498.
CrossRef |

Glasby P, Selkirk PM, Adamson D, Downing AJ, Selkirk DR (1988) Blue Mountains ash (Eucalyptus oreades R.T.Baker) in the western Blue Mountains. Proceedings of the Linnean Society of New South Wales 110, 141–158.

Graham AW, Wallwork MA, Sedgley M (1998) Lignotuber bud development in Eucalyptus cinerea (F.Muell. ex Benth.). International Journal of Plant Sciences 159(6), 979–988.
CrossRef |

Grose RJ (1961) Some silvical characteristics and notes on the silviculture of alpine ash, Eucalyptus delegatensis R.T.Baker. In ‘Proceedings of the second world Eucalyptus conference, Sao Paulo, Brazil’.

Hall N , Johnston RD , Chippendale GM (1975) ‘Forest trees of Australia.’ (Australian Government Publishing Service: Canberra)

Hedberg O (1964) Features of afroalpine plant ecology. Acta Phytogeographica Suecica 49, 1–14.

Hill KD (1989) Mallee eucalypt communities: their classification and biogeography. In ‘Mediterranean landscapes in Australia’. (Eds JC Noble, RA Bradstock) (CSIRO Publishing: Melbourne)

Hill KD (1991) Eucalyptus. In ‘Flora of New South Wales. Vol. 2.’ (Ed. GJ Harden) pp. 76–142. (New South Wales University Press: Sydney)

Hill KD, Johnson LAS (1991) Systematic studies in the eucalypts—3. New taxa and combinations in Eucalyptus (Myrtaceae) Telopea 4(2), 223–267.

Hill KD, Johnson LAS (1992) Systematic studies in the eucalypts. 5. New taxa and combinations in Eucalyptus (Myrtaceae) in Western Australia. Telopea 4(4), 561–634.

Hopper SD (1979) Biogeographical aspects of speciation in the southwest Australian flora. Annual Review of Ecology and Systematics 10, 399–422.
CrossRef |

Hopper SD (1993) Eucalypts. In ‘Mountains of mystery’. (Eds C Thomson, G Hall, G Friend) pp. 71–83. (Department of Conservation and Land management: Perth)

Hopper SD, Wardell-Johnson G (2004) Eucalyptus virginea and E. relicta (Myrtaceae), two new rare forest trees from south-western Australia allied to E. lane-poolei, and a new phantom hybrid. Nuytsia 15(2), 227–240.

Jacobs MR (1955) ‘Growth habits of the eucalypts.’ (Forestry and Timber Bureau: Canberra)

Johnson LAS, Hill KD (1991) Systematic studies in the eucalypts—2. A revision of the gimlets and related species: Eucalyptus extracodical series Salubres and Annulatae (Myrtaceae). Telopea 4(2), 201–222.

Karschon R (1967) Ecotypic variation in Eucalyptus camaldulensis Dehn. Contributions on Eucalypts in Israel 3, 35–53.

Kelly AE, Napier AC, Hopper SD (1995) Survey of rare and poorly known eucalypts of Western Australia. CALMScience (Suppl. 2), 1–207.

Kennington WJ, James SH (1997) Contrasting patterns of clonality in two closely related mallee species from Western Australia, Eucalyptus argutifolia and E. obtusiflora (Myrtaceae). Australian Journal of Botany 45, 679–689.
CrossRef |

Lacey CJ (1983) Development of large plate-like lignotubers in Eucalyptus botryoides Sm. in relation to environmental factors. Australian Journal of Botany 31, 105–118.
CrossRef |

Lacey CJ, Johnston RD (1990) Woody clumps and clumpwoods. Australian Journal of Botany 38, 299–334.
CrossRef |

Ladiges PY, Whiffin T (1993) Taxonomic revision of Eucalyptus alpina s.l. and recognition of three new species, E. victoriana, E. serraensis and E. verrucosa.  Australian Systematic Botany 6, 365–370.
CrossRef |

Ladiges PY, Humphries CJ, Brooker MIH (1987) Cladistic and biogeographic analysis of Western Australian species of Eucalyptus L’Herit., informal subgenus Monocalyptus Pryor & Johnson. Australian Journal of Botany 35, 251–281.
CrossRef |

Nicolle D, Conran JG (1999) Variation in the Eucalyptus flocktoniae complex (Myrtaceae) and the description of four new taxa from southern Australia. Australian Systematic Botany 12(2), 207–239.
CrossRef |

Nicolle D (2000a) A review of Eucalyptus calycogona (Myrtaceae) including the description of three new taxa from southern Australia. Nuytsia 13, 303–316.

Nicolle D (2000 b) ‘Currency Creek Arboretum eucalypt research. Vol. 1.’ (D. Nicolle: Adelaide)

Nicolle D (2002) Two new species of silver mallet (Eucalyptus—Myrtaceae) of very restricted distribution in south-western Western Australia. Nuytsia 15, 77–84.

Nicolle D (2003) ‘Currency Creek Arboretum eucalypt research. Vol. 2.’ (D. Nicolle: Adelaide)

Noble IR, Slatyer RO (1980) The use of vital attributes to predict successional changes in plant communities subject to recurrent disturbances. Vegetatio 43, 5–21.
CrossRef |

Prober SM, Austin MP (1990) Habitat peculiarity as a cause of rarity in Eucalyptus paliformis.  Australian Journal of Ecology 16, 189–205.

Prober SM, Bell JC, Moran GF (1990) A phylogenetic and allozyme approach to understanding rarity in three ‘green ash’ eucalypts. Plant Systematics and Evolution 172, 99–118.
CrossRef |

Pryor LD , Johnson LAS (1971) ‘A classification of the eucalypts.’ (Australian National University Press: Canberra)

Poynton RJ (1979) ‘Tree planting in southern Africa. Vol. 2. The eucalypts.’ (Department of Forestry, Republic of South Africa, Pretoria)

Rayner ME (1992) Application of dendrochronology, stem analysis and inventory data in the estimation of tree and stand ages in karri forest. Technical Report No. 27. Department of Conservation and Land Management, Perth.

Rossetto M, Jezierski G, Hopper SD, Dixon KW (1999) Conservation genetics and clonality in two critically endangered eucalypts from the highly endemic south-western Australian flora. Biological Conservation 88(3), 321–332.
CrossRef |

Smith S, Hughes J, Wardell-Johnson G (2003) High population differentiation and extensive clonality in a rare mallee eucalypt: Eucalyptus curtisii.  Conservation Genetics 4(3), 289–300.
CrossRef |

Specht RL (1970) Vegetation. In ‘The Australian environment’. 4th edn. (Ed. GW Leeper) pp. 44–67. (Melbourne University Press: Melbourne)

Tomlinson PB (1983) Structural elements of the rainforest. In ‘Tropical rain forest ecosystems. A. Structure and function’. (Ed. FB Golley) pp. 9–28. (Elsevier: Amsterdam)

Turnbull JW (1980) Provenance variation in Eucalyptus cloeziana. IUFRO conference on fast growing trees, Brazil.

Turnbull JW , Pryor LD (1984) Choice of species and seed sources. In ‘Eucalypts for wood production’. (Eds WE Hillis, AG Brown) pp. 6–65. (CSIRO and Academic Press: Sydney)

Tyson M, Vaillancourt RE, Reid JB (1998) Determination of clone size and age in a mallee eucalypt using RAPDs. Australian Journal of Botany 46, 161–172.
CrossRef |

Vaillancourt RE, Boomsma DB, Nicolle D (2001) An disjunct population of Eucalyptus globulus ssp. bicostata from South Australia. Transactions of the Royal Society of South Australia 125(1), 65–68.

Wardell-Johnson GW (2000) Response of forest eucalypts to moderate and high intensity fire in the Tingle Mosaic, south-western Australia: comparisons between locally endemic and regionally distributed species. Austral Ecology 25, 409–421.
CrossRef |

Wardell-Johnson G , Coates D (1996) Links to the past: local endemism in four species of forest eucalypts in southwestern Australia. In ‘Gondwanan heritage: past, present and future of the Western Australian biota’. (Eds SD Hopper, J Chappill, M Harvey, A George) pp. 137–154. (Surrey Beatty: Sydney)

Wardell-Johnson GW , Williams JE , Hill KD , Cumming R (1997) Evolutionary biogeography and contemporary distribution of eucalypts. In ‘Eucalypt ecology’. (Eds JE Williams, JCZ Woinarski) pp. 92–128. (Cambridge University Press: Cambridge, UK)

Western Australian Herbarium (2000) FloraBase—Information on the Western Australian flora. Department of Conservation and Land[verified 5 May 2006].

Whittock S, Steane DA, Vaillancourt RE, Potts BM (2003) Molecular evidence shows that the tropical boxes (Eucalyptus subgenus Minutifructus) are over-ranked. Transactions of the Royal Society of South Australia 127(1), 27–32.

Williams JE , Brooker MIH (1997) Eucalypts: an introduction. In ‘Eucalypt ecology’. (Eds JE Williams, JCZ Woinarski) pp. 1–15. (Cambridge University Press: Cambridge, UK)

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